In Madagascar's rainforests, we lack knowledge of the natural history of mistletoe, a hemiparasitic plant that is an essential food source and habitat for many animals. Here, we compared the germination success and survival of Bakerella gonoclada mistletoe seeds that were defecated by lemurs with seeds hand-removed from fruits to assess the effects of lemur gut passage on germination. We found that mistletoe seeds defecated by lemurs had significantly higher germination success, germination speed, hypocotyl growth, and survival probability than hand-removed seeds, demonstrating that lemurs enhance the germination of this species. These results suggest a mutualistic relationship between mistletoe and lemurs.

The movement decisions of frugivores shape spatial patterns of seed deposition, with considerable impacts for plant individuals, populations and communities. However, we have limited understanding of the mechanisms determining the foraging strategies behind the movements of frugivorous animals and their interaction with the landscape. Using spatially explicit simulations we examined the movements and seed dispersal of red-bellied lemur (Eulemur rubriventer) groups dispersing seeds of the tropical tree Harungana madagascariensis under four theorized foraging strategies. These included scenarios in which frugivores move directly to the (1) most rewarding plant in the landscape, (2) nearest fruiting plant, or towards the most rewarding plant in the landscape, with stops at any fruiting plants encountered along the way according to a local attraction distance of (3) 10 m and (4) 20 m. We then compared 12 metrics and data distributions for seed dispersal and movement patterns from these models to empirical observations. Simulations of foraging strategies with directed movement towards highly rewarding resources (strategy 1) produced the longest seed dispersal distances and best matched observed patterns. Observed data of lemur movements and directionality best matched results from three strategies, simulations towards highly rewarding resources without stops (strategy 1) and with either stops along the way with a 10 m (strategy 3) or 20 m (strategy 4) attraction distance. Collectively, our results supported the strategies in which lemurs moved consistently towards the small number of trees that produced the most fruit in the landscape. Our work supports incorporating a mechanistic understanding of animal foraging, movement behaviour and landscape patterns in fruit production for the prediction of seed movement. Explicit considerations of these mechanisms can improve our understanding of the maintenance and structure of diversity in plant communities, with implications for the conservation of forest ecosystems relying on animals for seed dispersal.Read the free Plain Language Summary for this article on the Journal blog.

Anthropogenic pressures such as hunting are increasingly driving the localized functional extinctions of all or most large and medium-sized wildlife species in tropical forests, a phenomenon broadly termed defaunation. Concurrently in these areas, smaller-bodied wildlife species benefit from factors such as competitive release and experience population increases. This transformation of the wildlife community can impact species interactions and ecosystem services such as seed dispersal and seed-mediated geneflow with far reaching consequences. Evidence for negative genetic effects following defaunation is well-documented in large-seeded plants that require large frugivores for long distance seed dispersal. However, how defaunation affects small-seeded (< 1.5cm diameter) plants, which are dispersed by frugivores with a wide range of body-sizes and responses to anthropogenic threats, is not well understood. To better understand the reach of defaunation’s impacts on tropical plant communities, we investigated spatial and genetic patterns in a hyperabundant small-seeded palm, Euterpe precatoria in three sites representing distinct defaunation levels. We found significantly higher fine-scale spatial genetic structure among nearest-neighbor seedlings in the defaunated site and in the recovering, partially defaunated site relative to the faunally-intact site. Defaunation was associated with shorter distances between seedlings and adults and lower genetic distance between adult and seedling cohorts. No effects were detected on inbreeding and genetic diversity; however, we caution that trends we detected indicate that defaunation influences the spatial distribution of genetic variation even in small-seeded plants that inherently have a broad suite of seed dispersal agents, and this could lead to negative downstream effects on genetic diversity.

Anthropogenic pressures such as hunting are increasingly driving the localised functional extinctions of large- and medium-sized wildlife in tropical forests, a phenomenon broadly termed ‘defaunation’. Concurrently in these areas, smaller-bodied species benefit from factors such as competitive release and increase in numbers. This transformation of the wildlife community can impact species interactions and ecosystem services such as seed dispersal and seed-mediated geneflow with far-reaching consequences. Evidence for negative genetic effects following defaunation is well-documented in large-seeded plants that require large frugivores for long-distance seed dispersal. However, how defaunation affects plants with small or medium-small seeds (< 1.5 cm), which tend to be consumed and dispersed by frugivorous mutualists of a range of body sizes and responses to anthropogenic threats, is not well understood. To better understand defaunation's impacts on tropical plant communities, we investigated spatial and genetic patterns in a hyperabundant medium-to-small-seeded palm, Euterpe precatoria in three sites with different defaunation levels. Results indicate that defaunation is associated with higher fine-scale spatial genetic structure among seedlings and increased spatial clustering within seedling cohorts and between seedlings and conspecific adults, as well as a reduction in nearest-neighbour distances between seedlings and conspecific adults. There were no clear effects on inbreeding or genetic diversity. However, we caution these trends may indicate that defaunation reduces seed dispersal services for species previously presumed to be robust to deleterious effects of losing large frugivores by virtue of having smaller seeds and broad suites of dispersal agents, and negative downstream effects on genetic diversity could occur.

While plant–animal interactions occur fundamentally at the individual level, the bulk of research examining the mechanisms that drive interaction patterns has focused on species or population levels. In seed‐dispersal mutualisms between frugivores and plants, little is known about the role of space and individual‐level variation among plants in structuring patterns of frugivory and seed dispersal in a plant community. Here we use a zoocentric approach to examine how space and variation between individual plants affect movement and visitation by frugivores foraging on individual fruiting plants. To do this, we used a spatially explicit network approach informed by observations of the movement and foraging of a frugivorous lemur species Eulemur rubriventer among individual plants in a diverse plant community in Madagascar. The resulting hierarchical networks show a few individual plants receiving the bulk of the interactions, demonstrating that a generalist frugivore species could act as an individual‐plant specialist within a plant community. The few individual plants that dominated interactions with lemurs shaped the modular spatial structure of frugivory interactions in the community and facilitated visitation to near neighbors. This interaction structure was primarily driven by extrinsic factors, as lemur movements among plants were significantly influenced by the individual plant's spatial position and the fruiting plant richness in its immediate neighborhood. Individual plants in central spatial locations with large fruit crops received the most visits. The observed inequality in the interactions of a generalist frugivore within a highly diverse plant community highlights the importance of considering individual‐level variation for essential ecosystem processes such as seed dispersal.

As environmental change and degradation accelerate, perturbing insect populations, we need to better understand the resource use dynamics of diverse wild pollinators. Most tropical trees are adapted for biotic pollination, yet we still know little about plant–pollinator interactions in African rainforests. We addressed this gap from a community perspective, identifying what floral traits – colour, scent, reward accessibility and visibility – structure visitation patterns among insect functional groups to tree species flowering in the understory of Korup National Park, Cameroon. To understand how visitor groups share resources, we used joint modelling that explicitly considers zero‐inflation in visitation rates and correlation among visitor groups. Most tree species had exposed floral rewards, and all were visited by multiple insect groups among which ants, bees, beetles, and flies were most abundant. Visitation rates varied more among tree species than among individual trees. Floral scent differences were important for structuring visits, particularly for flies, bees, and ants. Ant and bee visitation rates decreased while fly visits increased marginally throughout the season, correlated with the dry to wet season transition. Comparison with other lowland tropical understories suggests flies may be uniquely diverse and important to this system, and differences in seasonality and forest structure may be drivers of community differences. Floral scent is likely a key functional trait structuring flower–insect interactions in tropical forest environments and should be emphasised in future studies. Lastly, a joint modelling approach can elucidate community structure, particularly in communities with ecologically generalised interactions.

Edge effects, driven by human modification of landscapes, can have critical impacts on ecological processes such as species interactions, with cascading impacts on biodiversity as a whole. Characterizing how edges affect vital biotic interactions such as seed dispersal by frugivores is important for better understanding potential mechanisms that drive species coexistence and diversity within a plant community. Here, we investigated how differences between frugivore communities at the forest edge and interior habitats of a diverse tropical rainforest relate to patterns of animal-mediated seed dispersal and early seedling recruitment. We found that the lemur communities across the forest edge-interior gradient in this system showed the highest species richness and variability in body sizes at intermediate distances; the community of birds showed the opposite pattern for species richness. Three large-bodied frugivores, known to be effective dispersers of large seeds, tended to avoid the forest edge. As result, the forest edges received a lower rate of animal-mediated seed dispersal compared to the interior habitats. In addition, we also found that the seeds that were actively dispersed by animals in forest edge habitats were smaller in size than seeds dispersed in the forest interior. This pattern was found despite a similarity in seed size of seasonally fruiting adult trees and shrubs between the two habitats. Despite these differences in dispersal patterns, we did not observe any differences in the rates of seedling recruitment or seed-size distribution of successful recruit species. Our results suggest that a small number of frugivores may act as a potential biotic filter, acting on seed size, for the arrival of certain plant species to edge habitats, but other factors may be more important for driving recruitment patterns, at least in the short term. Further research is needed to better understand the potential long-term impacts of altered dispersal regimes relative to other environmental factors on the successional dynamics of edge communities. Our findings are important for understanding potential ecological drivers of tree community changes in forest edges and have implications for conservation management and restoration of large-seeded tree species in disturbed habitats.

Through frugivory and seed dispersal, vertebrates influence plant demography and forest regeneration. Variation in local habitat surrounding fruiting plants can influence frugivore foraging decisions, thereby creating intraspecific variation in seed dispersal services. However, we have little knowledge of drivers of local variation in frugivory. Here, we investigate factors that may influence frugivore diversity and fruit removal at the level of individual plants. We focus on a common understory palm within a continuous Choco forest with mixed land-use histories in Ecuador. The density of pioneer tree species in the genus Cecropia around focal palms was negatively related to fruit removal and the diversity of frugivores visiting palms. This may relate to the fact that the presence and abundance of Cecropia species often indicate the existence and severity of past disturbances. Local Cecropia density was also related to an overall shift in the frugivore community that corresponded with an increase in fruit removal by lower-quality seed dispersers (rodents). We also found that the local density of fruiting conspecifics was positively related to frugivore diversity, but not fruit removal. Our results provide information on drivers of intraspecific inequalities in plant populations across tropical forest landscapes. The reduction in fruit removal and frugivore diversity associated with local Cecropia abundance suggests that seed dispersal services can be sensitive to fine-scale variation in habitat structure. Furthermore, because Cecropia are often indicative of past disturbances, this indicates that even small-scale habitat degradation by humans can have lasting effects by creating localized pockets of forest unfavored by frugivores. Abstract in Spanish is available with online material.

ABSTRACT The relative abundance patterns of tropical trees have been of interest since the expeditions of Alfred Russel Wallace, but little is known about how differences in relative abundance relate to reproductive patterns. Flowering is resource-dependent and fitness differences as well as differences in the quality of the abiotic and biotic neighborhood may contribute to the variation in reproductive status responsible for population-level flowering patterns. This variation determines the density and distance between flowering conspecifics and may alter relative abundance extremes among species during reproduction, factors known to influence pollination success. We collected flowering status data for a guild of twenty-three co-occurring tree species that flower in the understory of the Korup Forest Dynamics Plot in Cameroon. We examined how the occurrence and location of reproductive events were related to spatial patterns of adult abundance, focal tree size, neighborhood crowding, and habitat, while accounting for the influence of shared ancestry. Across species, the probability of flowering was higher for individuals of rarer species and for larger individuals but was unrelated to neighborhood crowding or habitat differences. Relative abundance extremes were reduced when only flowering individuals were considered, leading to a negative relationship between plot abundance and flowering probability at the species level that was not structured by shared ancestry. Spatially, flowering conspecifics tended to be overdispersed relative to all adult conspecifics. Rare species are predicted to suffer Allee effects or reduced fitness due to the difficulty of finding mates at low densities and frequencies. Here, however, rare species appear to maximize the size of their mate pool, compared to abundant species. If this partial ‘leveling of the playing field’ during reproduction is typical, it has consequences for our understanding of biodiversity maintenance and species coexistence in tropical forests.

Throughout the world, biodiversity and nature's contributions to people are under threat, with clear changes evident. Biodiversity and ecosystem services have particular value in Africa– yet they are negatively impacted by a range of drivers, including land use and climate change. In this communication, we show evidence of changing biodiversity and ecosystem services in Africa, as well as the current most significant drivers of change. We then consider five plausible futures for the African continent, each underlain by differing assumptions. In three out of the five futures under consideration, negative impacts on biodiversity and ecosystem services are likely to persist. Those two plausible futures prioritizing environment and sustainability, however, are shown as the most likely paths to achieving long term development objectives without compromising the continent's biodiversity and ecosystem services. Such a finding shows clearly that achievement of such objectives cannot be separated from full recognition of the value of such services.

With 60% of all primate species now threatened with extinction and many species only persisting in small populations in forest fragments, conservation action is urgently needed. But what type of action? Here we argue that restoration of primate habitat will be an essential component of strategies aimed at conserving primates and preventing the extinctions that may occur before the end of the century and propose that primates can act as flagship species for restoration efforts. To do this we gathered a team of academics from around the world with experience in restoration so that we could provide examples of why primate restoration ecology is needed, outline how primates can act as flagship species for restoration efforts of tropical forest, review what little is known about how primate populations respond to restoration efforts, and make specific recommendations of the next steps needed to make restoration of primate populations successful. We set four priorities: (1) academics must effectively communicate both the value of primates and the need for restoration; (2) more research is needed on how primates contribute to forest restoration; (3) more effort must be put into Masters and PhD level training for tropical country nationals; and finally (4) more emphasis is needed to monitor the responses of regenerating forest and primate populations where restoration efforts are initiated. We are optimistic that populations of many threatened species can recover, and extinctions can be prevented, but only if concerted large-scale efforts are made soon and if these efforts include primate habitat restoration.

Hunting has reduced or eliminated large-bodied vertebrates in many areas across the tropics, contributing to the global process of defaunation. Elucidating the ecological consequences of hunting has important implications for managing ecosystems and for our understanding of community and ecosystem ecology. We present data collected through a combination of comparative and experimental approaches to assess how faunally-intact and heavily-hunted forests in Gabon differ in understory vegetation structure, macroinvertebrate fauna, ecological processes, and the relative importance of different taxa driving those processes. Our results show that hunted sites had denser understory vegetation and hosted approximately 170 times fewer termites compared to faunally-intact sites. While web-building spiders were positively associated with understory vegetation density, this effect did not translate to significantly higher abundances in heavily-hunted forests. Additionally, the overall rates of decomposition, insectivory, and seed predation/removal on the forest floor appeared robust to both defaunation and the associated increases in understory vegetation density. However, our exclosure experiments revealed that the contribution of invertebrates to decomposition was approximately 25% lower in hunted sites compared to faunally-intact sites. Results suggest potential resilience in this complex ecosystem such that microbial or other taxa not measured in this study may compensate for the reduced functional contribution of invertebrates to decomposition. However, while our results illustrate potential resilience, they also indicate that indirect effects following defaunation, such as increases in the density of understory vegetation, may alter invertebrate communities on the forest floor, with potential consequences for the mechanisms, and therefore the dynamics, driving critical ecosystem processes.

As the influence of climate change on tropical forests becomes apparent, more studies are needed to understand how changes in climatic variables such as rainfall are likely to affect tree phenology. Using a twelve‐year dataset (2005–2016), we studied the impact of seasonal rainfall patterns on the fruiting phenology of 69 tree species in the rain forest of southeastern Madagascar. We found that average annual rainfall in this region has increased by >800 mm (23%) during this period relative to that recorded for the previous 40 years and was highly variable both within and between years. Higher monthly measures of fruiting richness and the intensity of fruiting in our sample community were associated with significantly higher levels of rainfall. We also found that less rainfall during the dry season, but not the wet season, was associated with a significant shift toward later timing of peak richness and peak intensity of fruiting in the subsequent 12 months; however, this pattern was driven primarily by an extreme drought event that occurred during the study period. Longer time scales of phenology data are needed to see whether this pattern is consistent. Madagascar is expected to experience more extremes in rainfall and drought with increasing climate change. Thus, the linkages between variable precipitation and the fruiting phenology of forest trees will have important consequences for understanding plant reproduction and the ability of Madagascar's wildlife to cope with a changing climate.

Primate seed dispersal is a vital, but complex, ecological process that involves many interacting agents and plays important roles in the maintenance of old-growth forest, as well as in the development of regenerating forest. Focusing primarily on African examples, in this article we briefly review the ecological process of primate seed dispersal, highlighting understudied and contentious topics, and then we discuss how our knowledge on primate seed dispersal can promote both forest restoration and primate conservation. Though it is frequently claimed that primates are critically important for the maintenance of diverse tropical forest ecosystems, we believe that more empirical evidence is needed to support this claim. Confounding factors can often be difficult to rule out and long-term studies extending beyond the seedling or sapling stage are very rare. In addition, though primates are critical for initial seed dispersal of many tree species, spatial and temporal variation in post-deposition processes, such as secondary seed dispersal and predation by rodents, can dramatically alter the initial patterns generated by primates. However, given the need for immediate conservation action to prevent further primate extinctions, we advocate that the knowledge about primate seed dispersal be used in formulating informed conservation plans. One prominent area where this knowledge will prove extremely valuable is in forest restoration efforts. To aid in the development of such efforts, we pose five questions, the answers to which will help facilitate forest restoration becoming a useful tool in strategies designed to conserve primates.

1. Edge effects are among the most significant consequences of forest fragmentation. Therefore, understanding the impacts of edge creation on biodiversity is crucial for forest management and biological conservation. 2. In this study, we used trait-based and phylogenetic approaches to examine the effects of fragmentation on components of diversity and above-ground biomass of rainforest tree communities in Madagascar in forest edge vs. interior habitats. 3. Tree communities in forest edges showed lower phylogenetic diversity relative to those in interior habitats, suggesting that some clades may be more vulnerable to environmental filtering than others. Functional diversity was also significantly lower on the edge for productivity traits, but not for dispersal traits. Tree communities in the forest edge showed higher divergence of dispersal traits and lower divergence in productivity traits than expected, while functional diversity in interior forest did not differ from random expectations. This suggests that separate mechanisms affect productivity traits vs. dispersal traits in edge habitats. 4. There was no significant difference in above-ground biomass between edge and interior habitats, suggesting that edge effects have not yet negatively influenced the forest's potential for carbon storage. However, these changes may not have occurred yet, given the slow turnover of tree communities. 5. Synthesis and applications. Our results highlight the role of edge effects in the erosion of functional and phylogenetic diversity of highly diverse tree communities. While above-ground biomass did not appear to be affected by forest edge in our study, we suggest long-term monitoring of forests for potential changes in ecosystem functioning. These findings also indicate the need to reduce edge creation and buffer existing edges for holistic biodiversity conservation.

Anthropogenic disturbances have resulted in declines of seed-dispersing primate frugivores in tropical forests. Previous work has suggested that loss of seed dispersal by large frugivores may have a negative impact on ecosystem carbon storage by reducing tree biomass. However, we know little about the potential impacts of losing frugivores in Madagascar’s diverse rainforest ecosystem. Understanding the effects of frugivore extinction on carbon loss is relevant in Madagascar, where threatened lemur taxa are the only dispersers of many large-seeded plant species. Using a dataset of tree species composition and traits from the southeastern rainforests of Ranomafana National Park, we examined whether seed size and lemur-dependent dispersal are positively associated with above-ground tree biomass. We then simulated different scenarios of population declines of large-seeded trees (>10 mm seed length) dependent on lemur-mediated seed dispersal, to examine potential directional changes in carbon storage capacity of Malagasy forests under lemur loss. Lemur-dispersed tree species, which have large seeds, had higher above-ground biomass than other species. Our simulations showed that the loss of large frugivorous primates in Madagascar may decrease the forest’s potential to store carbon. These results demonstrate the importance of primate conservation for maintaining functioning ecosystems and forest carbon stocks in one of the world’s hottest hotspots of biodiversity.